Diversity and host recurrence of fungi associated with the bleached leaf litter in a subtropical forest

Fungi that selectively remove lignin cause extensive bleaching of leaf litter, which results in the acceleration of litter decomposition. The taxonomic diversity and host recurrence of saprotrophic fungi associated with bleached leaf litter were investigated in a subtropical forest in southern Japan. A total of 211 fungal isolates and sporocarps were obtained from bleached portions of leaf litter of 33 plant species in 18 families and were classified into 83 operational taxonomic units (OTUs) at the 97% similarity level of the ITS rDNA sequence. These fungal OTUs belonged to Rhytismataceae and Xylariaceae in the Ascomycota, and Marasmiaceae, Mycenaceae, Lachnocladiaceae, and Physalacriaceae in the Basidiomycota. OTUs in Rhytismataceae and Marasmiaceae showed a significantly higher degree of recurrence for plant species than simulated networks where partners were associated randomly. In contrast, OTUs in Xylariaceae and Mycenaceae showed no significant recurrence for plant species. Such differing degrees of recurrence for plant species implied different roles of fungal families in leaf litter decomposition.     

Wood decomposing abilities of diverse lignicolous fungi on nondecayed and decayed beech wood

We tested the decay abilities of 28 isolates from 28 lignicolous fungal species (Basidiomycota, Ascomycota and Zygomycota) with the pure culture test. We used beech wood powder in varying moisture conditions and decay stages (nondecayed, intermediately decayed and well decayed) as substrates. The weight loss in wood powder was -0.2-17.8%. Five isolates of Basidiomycota (Bjerkandera adusta, Mycena haematopus, Omphalotus guepiniformis, Trametes hirsuta, Trametes versicolor) caused high weight losses in nondecayed wood. We detected significant effects of decay stage on weight loss in wood in most isolates tested, whereas moisture content rarely had an effect on weight loss. Among Basidiomycota and Xylariaceae in Ascomycota weight loss was greater for nondecayed wood than for intermediately and well decayed wood. In contrast four isolates in Ascomycota (Scytalidium lignicola, Trichoderma hamatum, T. harzianum, T. koningii) caused substantial weight loss in intermediately and well decayed wood, although they rarely caused weight loss in nondecayed wood. Zygomycota caused low weight loss in wood. Wood decay stages also affected decomposition of wood chemical components. Acid-unhydrolyzable residue (AUR) decomposition was reduced, whereas holocellulose decomposition was stimulated by some strains of Basidiomycota and Ascomycota in well decayed wood. T. harzianum in particular caused significant weight loss of holocellulose in well decayed wood, although this fungus caused negligible weight loss of both AUR and holocellulose in nondecayed wood. We discuss these changes in the decay patterns of AUR and holocellulose with varying wood decay stages in relation to the role of fungal decomposition of woody debris in forests.     

Organic chemical and nutrient dynamics in decomposing beech leaf litter in relation to fungal ingrowth and succession during 3-year decomposition processes in a cool temperate deciduous forest in Japan

Decomposition processes of beech leaf litter were studied over a 3-year period in a cool temperate deciduous forest in Japan. Organic chemical and nutrient dynamics, fungal biomass and succession were followed on upper (Moder) and lower (Mull) of a forest slope. Litter decomposition rates were similar between the sites. Nutrient dynamics of the decomposing litter was categorized into two types: weight changes in nitrogen and phosphorus showed two phases, the immobilization (0–21 months) and the mobilization phase (21–35 months), while those in potassium, calcium and magnesium showed only the mobilization phase. The rate of loss of organic chemical constituents was lignin < holocellulose < soluble carbohydrate < polyphenol in order. The changes in lignocellulose index (LCI), the ratio of holocellulose in lignin and holocellulose, were significantly correlated to the changes in nitrogen and phosphorus concentrations during the decomposition. During the immobilization phase, increase in total fungal biomass contributed to the immobilization of nitrogen and phosphorus. The percentage of clamp-bearing fungal biomass (biomass of the Basidiomycota) to total fungal biomass increased as the decomposition proceeded and was significantly correlated to LCI. Two species in the xylariaceous Ascomycota were dominantly isolated by the surface sterilization method from decomposing litter collected in the 11th month. The organic chemical, nitrogen and phosphorus dynamics during the decomposition were suggested to be related to the ingrowth, substrate utilization and succession of the Xylariaceae and the Basidiomycota. Twenty-one species in the other Ascomycota and the Zygomycota isolated by the washing method were classified into three groups based on their occurrence patterns: primary saprophytes, litter inhabitants and secondary sugar fungi. These species showed different responses to LCI and soluble carbohydrate concentration of the litter between the groups.     

Roles of diverse fungi in larch needle-litter decomposition

Functional biodiversity of fungi in larch (Larix leptolepis) forests needle-litter decomposition was examined by a pure-culture test. Weight loss of larch-needle litter, utilization pattern of lignocellulose and chemical composition of remaining litter were investigated and compared for 31 isolates in 27 species of basidiomycetes and ascomycetes. Weight loss (% original weight) of litter ranged from -2.0% to 14.2%. Mean weight loss of litter caused by the basidiomycetes was not significantly different from that caused by the ascomycetes. Basidiomycetes caused loss of lignin and carbohydrates in variable proportions, while ascomycetes exclusively attacked carbohydrates without delignification. The content of lignin and nitrogen in remaining litter was not significantly correlated when both basidiomycetes and ascomycetes were included. However, the correlation coefficient was significant when the relationship was examined separately for basidiomycetes, indicating that the degree of selective delignification determined the final nitrogen content in litter. Possible effects of fungal colonization on needle-litter decomposition in larch forests are discussed.     

Fungal decomposition of Abies needle and Betula leaf litter

The effect of litter type and incubation temperature on the ability of fungi to decompose leaf litter of subalpine trees was examined by a pure-culture test. Mass loss of Abies needle and Betula leaf litter and utilization patterns of lignin and carbohydrates were investigated under two temperature conditions (20 C and 10 C) and compared for 29 species in basidiomycetes, ascomycetes and zygomycetes. The decomposing ability was generally higher in basidiomycetes than in ascomycetes and zygomycetes. Mass loss (% original mass) of litter was higher in Betula than in Abies and higher at 20 C than at 10 C. The 29 fungi were divided into lignocellulose decomposers, cellulose decomposers and sugar fungi based on their substrate utilization in Abies and Betula litter. Mass loss of lignin and carbohydrates by lignocellulose and cellulose decomposers was higher in Betula than in Abies. Mass loss of carbohydrates was higher at 20 C than at 10 C, but the temperature did not influence mass loss of lignin, indicating lignin decomposition by fungi was less sensitive to temperature than carbohydrate decomposition. Lignin/carbohydrate loss ratio (L/C) of Collybia spp. that caused selective delignification was lower at 20 C than at 10 C. These results indicate that the decomposability of litter, lignin and carbohydrate was different between Abies and Betula and that temperature affected not only the rate at which fungi decompose litter but also the ability of fungi to use lignin and carbohydrates.     

A literature synthesis resolves litter intrinsic constraints on fungal dynamics and decomposition across standing dead macrophytes

The standing dead phase is an important stage in the decomposition of emergent vegetation in marsh wetlands, yet few studies have examined how intrinsic litter traits constrain rates of standing litter decomposition or fungal colonization across plant tissue types or species. To address broad constraints on the decomposition of standing dead litter, we conducted a systematic survey of emergent standing dead decomposition studies that measured decay rates and/or fungal biomass, and litter % lignin, carbon:nitrogen (C:N) and/or carbon:phosphorus (C:P). Across 52 datasets, litter of low C:N and C:P ratios exhibited increased decomposition rates (r = −0.737 and −0.645, respectively), whereas % lignin was not significantly correlated with decomposition rates (r = 0.149). Mixed-effects models for litter decomposition rates indicated significant effects of litter molar C:N and C:N + lignin as an additive model, with the former providing marginally better support. Litter % lignin, however, was strongly negatively correlated with fungal biomass (r = −0.669), indicating greater fungal colonization of low-lignin litter, and not correlated with C:N (r = −0.337) and C:P (r = −0.456) ratios. The best-supported model predicting fungal biomass was litter molar C:N, with the C:N + lignin additive model also showing significant effects. Fungal carbon-use efficiency (CUE) also had a strong negative correlation with % lignin (r = −0.604), molar C:N (r = −0.323) and C:P (r = −0.632) across datasets. Our study demonstrates the constraining effects that litter stoichiometry and % lignin elicit on decomposition of standing dead litter and fungal colonization, respectively. These findings improve our understanding of biogeochemical cycling and prediction of the fates of C and nutrients in wetlands.     

Litter Decomposition in a California Annual Grassland: Interactions Between Photodegradation and Litter Layer Thickness

In annual grasslands that experience a mediterranean-type climate, the synchrony between plant senescence and peak solar radiation over summer results in high litter sun exposure. We examined the decomposition of both shaded and sun-exposed litter over summer and inferred the effects of photodegradation from changes in mass loss and litter chemistry. The carry-over effects of summer litter exposure on wet season decomposition were also assessed, and the attenuation of photodegradation with litter layer thickness was used to estimate the proportion of grass litter lignin susceptible to photodegradation under different treatments of a factorial global change experiment. Over summer, mass loss from grass and forb litter exposed to ambient sunlight ranged from 8% to 10%, whereas lignin decreased in grass litter by approximately 20%. After one year of decomposition, mass losses from grass leaves exposed to sunlight over summer were more than double the mass losses from summer-shaded leaves. When shade litter layer thickness was varied, mass losses over summer for all treatments were also approximately 8%; however, lignin decreased significantly only in the low shade treatments (0–64 g m⁻² of shade litter). Aboveground production of annual grasses nearly quadrupled in response to the combined effects of N addition, elevated atmospheric CO₂, increased precipitation and warming. The estimated proportion of grass litter lignin experiencing full photodegradation ranged from 100% under ambient conditions to 31–62% in plots receiving the combined global change treatments. These results reveal an important role of sun exposure over summer in accelerating litter decomposition in these grasslands and provide evidence that future changes in the quantity of litter deposition may modulate the influence of photodegradation integrated across the litter layer.     

人工调控措施下马尾松凋落叶化学质量变化及与分解速率的关系

马尾松凋落叶分解缓慢,促进其凋落物分解,提高养分归还速度,维持地力稳定,已成为马尾松人工林可持续经营中的关键问题。基于此,采用正交试验L_9(3~4)设计,选择菌剂、表面活性剂、不同碳氮营养液和有机肥料4种人工调控因素,在马尾松林下开展凋落叶分解调控试验,以掌握不同调控组合对凋落叶分解速率和化学质量的影响及作用效果等。结果表明:有机肥料和菌剂显著影响马尾松凋落叶分解速率,腐解剂2和鸡粪联合作用更利于分解。马尾松凋落叶在林下自然分解过程中,化学质量参数向着利于分解的方向变化,N、P以积累为主,C/N、C/P、L/N和L/P呈降低态势,人为调控措施加速了这一变化进程;不同调控措施对凋落叶化学质量参数的影响不尽相同,添加有机肥料有利于剩余凋落叶N、P含量升高,C/N、C/P、L/N和L/P的降低;菌剂腐解剂2有利于L/P、C/P的降低;表面活性剂OP-10有利于凋落叶L/N的降低。人工调控下,调控因素可通过改变凋落物化学质量影响其分解速率,N含量和C/N是影响马尾松凋落叶分解速率的主要因素;而P浓度、L/N、C/P、L/P对分解速率的影响不规律或不显著。     

Decomposition of [C]Lignocelluloses of Spartina alterniflora and a Comparison with Field Experiments

Decomposition of lignocelluloses from Spartina alterniflora in salt-marsh sediments was measured by using C-labeled compounds. Rates of decomposition were fastest in the first 4 days of incubation and declined later. Lignins labeled in side chains were mineralized slightly faster than uniformly labeled lignins; 12% of the [side chain-C]lignin-labeled lignocellulose was mineralized after 816 h of incubation, whereas only 8% of the [U-C]lignin-labeled lignocelluloses were degraded during this period. The carbohydrate moiety within the lignocellulose complex was degraded about four times faster than the lignin moiety; after 816 h of incubation, 29 to 37% of the carbohydrate moiety had been mineralized. Changes in concentration of lignin and cellulose in litter of S. alterniflora were followed over 2 years of decay. Cellulose disappeared from litter more rapidly than lignin; 50% of the initial content of cellulose was lost after 130 days, whereas lignin required 330 to 380 days for 50% loss. The slow loss of lignin compared with other litter components resulted in a progressive enrichment of litter in lignin content. The rates of mineralization of [C]lignocelluloses in marsh sediments were similar to the rates of lignocellulose decomposition in litter on the marsh.     

川西高山林线交错带凋落叶分解速率与初始质量的关系

对我国川西高山林线交错带14种代表性植物凋落叶分解速率与初始质量的关系进行研究.结果表明: 高山林线交错带植物凋落叶分解速率（k）为0.16～1.70,乔木和苔藓凋落叶分解较慢,灌木凋落叶次之,草本凋落叶分解最快.凋落叶分解速率与N、木质素、酚类物质、C/N、C/P、木质素/N均具有显著的线性回归关系.通径分析得出,木质素/N和半纤维素含量可以解释k变异的78.4%,其中木质素/N可以解释k变异的69.5%,木质素/N对k的直接通径系数为-0.913.主成分分析表明,第1排序轴k、分解时间（t）的贡献率达99.2%,木质素/N、木质素含量、C/N、C/P与第1排序轴呈显著正相关,其中木质素/N与第1排序轴的相关关系最强(r=0.923).木质素/N是影响川西高山林线交错带植物凋落叶分解速率的关键质量指标,且凋落叶初始木质素/N越高,分解速率越低.     

Irrigation,fertilization and initial substrate quality effects on decomposing Loblolly pine litter chemistry

Changes in carbon chemistry (i.e., carbon compound classes such as aromatics, phenolics, etc.) of loblolly pine (Pinus taeda L.) litter were examined during three years of decomposition under factorial combinations of irrigation and fertilization treatments. Cross polarization magic angle spinning ¹³C nuclear magnetic resonance revealed that total carbon and nutrient concentrations correlated strongly with carbohydrate and O-alkyl carbon concentrations but did not relate well with concentrations of lignin, aromatic and phenolic carbon, or with lignin-related decomposition indices. The best correlations to carbon and nutrient concentrations occurred with the C/N (R²=0.86, P > 0.0001) and alkyl/O-alkyl (R²=0.75, P > 0.0001) decomposition indices. In all situations, the carbon chemistry of the decomposing litter followed the general pattern of accumulation of alkyl and carbonyl carbon with a loss of O-alkyl and methoxy carbon. Only small variations in the aromatic and phenolic carbon concentrations were detected. Since lignin is composed primarily of aromatic and phenolic carbons, the observation that there were only small changes in the aromatic and phenolic carbons of the litter is consistent with the general stability of lignin in these ecosystems. Trends in carbon chemistry during decomposition suggested that fertilization accelerated the decomposition process by about 100% as compared with the control plots. Irrigation also accelerated the decomposition process but to a lower extent (about 62% greater than control plots). Initial litter quality, as defined by the litter C/N, did not have a significant effect on the carbon chemistry of the decomposing litter. This study demonstrated that the decomposition mechanisms were not altered by the treatments but there were important changes in the relative chemistry of the decomposing litter which impacted the rate of decomposition.     

Lignin properties in topsoils of a beech/oak forest after 8 years of manipulated litter fall: relevance of altered input and oxidation of lignin

Background and aims

We studied the response of lignin oxidation in soils of a beech/oak forest to changes in litter fall. Additionally we considered possible factors in lignin oxidation, including altered (i) input of fresh organic matter and (ii) fungi-to-bacteria ratios.

Methods

The field-based experiment included (i) doubling and (ii) exclusion of litter fall and (iii) controls with ambient litter fall. Soil (0–20 cm depth) was sampled after 8 years. We analyzed (i) lignin using the CuO oxidation method, (ii) stocks of free and mineral-bound organic carbon (OC), (iii) the response of soil organic matter (SOM) decomposition to addition of labile organic compounds in laboratory incubations, and (iv) ratios of fungal- vs. bacterial-derived amino sugars (F/B ratios).

Results

Litter exclusion increased stocks of free-light fraction OC, F/B ratios, the ability of the microbial community to use labile compounds for SOM decomposition, as well as acid-to-aldehyde ratios of vanillyl-type lignin phenols in A horizons. Litter addition had no such effects. We assume that litter exclusion caused enhanced transport of organic debris from lower forest floor horizons with rainwater into the A horizon. Enhanced input of organic debris might have increased (i) the availability of labile compounds and (ii) F/B ratios. Consequently, lignin oxidation increased.

Conclusions

Enhanced input of organic debris from forest floors can increase lignin oxidation in mineral topsoils of the studied forest. The expected gradual changes in litter fall due to climate change likely will cause no such effects.     

中国森林生态系统凋落叶分解速率的分布特征及其控制因子

凋落物分解是森林生态系统碳循环的重要组成部分。建立中国森林凋落叶分解速率数据库, 分析凋落叶分解速率与其主要影响因素之间的关系, 对精确地预测中国森林生态系统碳收支具有重要意义。该研究通过收集已报道的中国森林凋落叶分解常数(k)及其相关变量, 分析探讨地理因素(纬度、经度和海拔)、气候因素(年平均气温和年降水量)、凋落叶质量(氮、磷、钾、木质素、木质素:氮和碳氮比)和叶特性(常绿与落叶、阔叶与针叶)对中国森林凋落叶分解速率的影响。结果表明, 在国家尺度上, k随年平均气温、年降水量、氮、磷和钾的增加而增加, 随纬度、经度、海拔、碳氮比、木质素和木质素:氮的增大而减小, 叶特性对k的影响不显著。气候与地理因素(年平均气温、年降水量和纬度)能解释k值变异的34.1%, 凋落叶质量(氮、钾、木质素和木质素:氮)能解释k值变异的21.7%, 它们能共同解释k值变异的74.4%。了解森林凋落叶分解速率在国家尺度上的格局和主控因素可为中国森林生态系统碳循环相关模型提供基础参数。     

Decomposition of [14C]Lignocelluloses of Spartina alterniflora and a Comparison with Field Experiments

Decomposition of lignocelluloses from Spartina alterniflora in salt-marsh sediments was measured by using ¹⁴C-labeled compounds. Rates of decomposition were fastest in the first 4 days of incubation and declined later. Lignins labeled in side chains were mineralized slightly faster than uniformly labeled lignins; 12% of the [side chain-¹⁴C]lignin-labeled lignocellulose was mineralized after 816 h of incubation, whereas only 8% of the [U-¹⁴C]lignin-labeled lignocelluloses were degraded during this period. The carbohydrate moiety within the lignocellulose complex was degraded about four times faster than the lignin moiety; after 816 h of incubation, 29 to 37% of the carbohydrate moiety had been mineralized. Changes in concentration of lignin and cellulose in litter of S. alterniflora were followed over 2 years of decay. Cellulose disappeared from litter more rapidly than lignin; 50% of the initial content of cellulose was lost after 130 days, whereas lignin required 330 to 380 days for 50% loss. The slow loss of lignin compared with other litter components resulted in a progressive enrichment of litter in lignin content. The rates of mineralization of [¹⁴C]lignocelluloses in marsh sediments were similar to the rates of lignocellulose decomposition in litter on the marsh.     

Litter decay rates are determined by lignin chemistry

Litter decay rates are often correlated with the initial lignin:N or lignin:cellulose content of litter, suggesting that interactions between lignin and more labile compounds are important controls over litter decomposition. The chemical composition of lignin may influence these interactions, if lignin physically or chemically protects labile components from microbial attack. We tested the effect of lignin chemical composition on litter decay in the field during a year-long litterbag study using the model system Arabidopsis thaliana. Three Arabidopsis plant types were used, including one with high amounts of guaiacyl-type lignin, one with high aldehyde- and p-hydroxyphenyl-type lignin, and a wild type control with high syringyl-type lignin. The high aldehyde litter lost significantly more mass than the other plant types, due to greater losses of cellulose, hemicellulose, and N. Aldehyde-rich lignins and p-hydroxyphenyl-type lignins have low levels of cross-linking between lignins and polysaccharides, supporting the hypothesis that chemical protection of labile polysaccharides and N is a mechanism by which lignin controls total litter decay rates. 2D NMR of litters showed that lignin losses were associated with the ratio of guaiacyl-to-p-hydroxyphenyl units in lignin, because these units polymerize to form different amounts of labile- and recalcitrant-linkages within the lignin polymer. Different controls over lignin decay and polysaccharide and N decay may explain why lignin:N and lignin:cellulose ratios can be better predictors of decay rates than lignin content alone.     

Lignocellulose degradation and crude protein formation by three ligninolytic Streptomyces strains

Streptomyces strains (11A, 21B and 31C) were grown on guinea grass lignocellulose and their ability to decompose lignocellulose was monitored. All three Streptomyces strains caused lignocellulose weight losses ranging from 27 to 34%. The three Streptomyces strains were also found to metabolize between 27 and 40% of the lignin component and 22–29% of the carbohydrate component of lignocellulose over the 12 week incubation period. Crude protein increases of degraded guinea grass lignocellulose were between 10–1 and 14–5% over the same period.     

Spatial characteristics in decomposition rate of foliar litter and controlling factors in Chinese forest ecosystems

凋落物分解是森林生态系统碳循环的重要组成部分。建立中国森林凋落叶分解速率数据库, 分析凋落叶分解速率与其主要影响因素之间的关系, 对精确地预测中国森林生态系统碳收支具有重要意义。该研究通过收集已报道的中国森林凋落叶分解常数(k)及其相关变量, 分析探讨地理因素(纬度、经度和海拔)、气候因素(年平均气温和年降水量)、凋落叶质量(氮、磷、钾、木质素、木质素:氮和碳氮比)和叶特性(常绿与落叶、阔叶与针叶)对中国森林凋落叶分解速率的影响。结果表明, 在国家尺度上, k随年平均气温、年降水量、氮、磷和钾的增加而增加, 随纬度、经度、海拔、碳氮比、木质素和木质素:氮的增大而减小, 叶特性对k的影响不显著。气候与地理因素(年平均气温、年降水量和纬度)能解释k值变异的34.1%, 凋落叶质量(氮、钾、木质素和木质素:氮)能解释k值变异的21.7%, 它们能共同解释k值变异的74.4%。了解森林凋落叶分解速率在国家尺度上的格局和主控因素可为中国森林生态系统碳循环相关模型提供基础参数。     

Effects of streams on lignin degradation during foliar litter decomposition in an alpine forest

AimsStreams are widely distributed in alpine forests, and litter decomposition in which is an important component of material cycling across the forest landscape. The leaching and fragmenting effects as well as the unique environmental factors in streams may have significant impacts on lignin degradation during litter decomposition, but studies on this are lacking.
Methods Using litterbag methods, we investigated the dynamics of lignin mass remaining and concentration (percent litter mass, %) during the decomposition of four foliar litters, which varied significantly in the initial litter chemical traits, from the dominant species of Salix paraplesia, Rhododendron lapponicum, Sabina saltuaria, and Larix mastersiana under different habitats (forest floor, stream, and riparian zone) in the upper reaches of the Minjiang River.
Important findings After two year’s incubation, litter lignin mass remaining for a specific litter species varied significantly (p < 0.05) among habitats, with an order of stream < riparian zone < forest floor. Lignin was degraded substantially in the early stage of litter decomposition process, and the lignin concentration first decreased and then increased with the proceeding of litter decomposition, but varied significantly (p < 0.05) among different litter species. Lignin mass showed a general trend of decrease across the 2-year decomposition course. In addition, habitat type, decomposition period and microenvironmental factors (e.g., temperature, pH value and nutrient availability) showed substantial influences on lignin degradation rate. These results suggest that the traditional view that lignin was relatively recalcitrant with an increase of concentration in the early stage of litter decomposition is challenged, but the loss of lignin in the early phrase is in line with recent findings about the fate of lignin during litter decomposition. Moreover, the significant differences of lignin degradation rates among different decomposition period and habitat types indicated that local-scale environmental factors can play a significant role in litter decomposition and lignin degradation processes.     

季节性冻融期间亚高山森林凋落物的质量变化

凋落物质量是影响凋落物分解的重要生物因子,其在季节性冻融期间的变化可能对亚高山森林生态系统过程产生显著的影响。因此,采用凋落物分解袋法,研究了岷江冷杉(Abies faxoniana)和白桦(Betula platyphylla)凋落物质量在一个季节性冻融期间(2006年10月至2007年4月)的变化。季节性冻融期间,岷江冷杉和白桦凋落物的木质素(L)和纤维素的降解率为全年降解的70%-75%,岷江冷杉和白桦凋落物的C/N、L/N和纤维素/N均显著增加,而纤维素/P均有所降低。岷江冷杉凋落物的C/P和L/P有所增加,但白桦凋落物的C/P和L/P有所降低。可见,季节性冻融期间,亚高山森林凋落物的质量发生了较为显著的变化,其显著影响了亚高山凋落物分解过程。